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-8 9 - 

A. J. Grout. 
There has been a considerable demand for a simple book on the 
Hepatics. To meet this demand I am preparing a treatment of the Hepaties 
similar to that which I have given the mosses in "Mosses with a Hand- 
Lens." This will be included in the second edition of that book now in pre- 
paration (See adv. in this No. of the Bryologist). This key to the genera is 
printed here with the hope that it will be used and criticised by the readers 
of the Bryologist, and that by the assistance of these criticisms the final 
treatment may be made more helpful. With the Queen ^-inch achromatic 
triplet I am able to make out the more minute structures mentioned in the 
keys. Many of them, especially leaf structure, can not be made out satisfac- 
torily unless the objects be mounted in water on a slide in the same man- 
ner as for a compound microscope. The slide should then be held up to the 
strong light, the slide being held with the left hand and the lens with the 
right, the right thumb resting upon that of the left hand so that the focus 
will not be distributed by any unsteadiness of the hands. 

From now until winter closes in I shall be glad to attempt to name 
Hepatics for our subscribers if the speciments be accompanied by a stamp, 
full data for the label, and the best name the collector can give. Fresh 
material only is desired. Almost none of the books give the time of matur- 
ing spores of the different species, and I hope that our readers will send me 
all the data of this sort that they have. Comparatively few illustrations are 
possible in this article, but the figures in the sixth edition of Gray's Manual 
will prove very helpful. In working up this key I have been surprised to 
find that sterile Hepatics are, as a rule, much easier to identify than sterile 
mosses. Many of the species maturing their spores in early spring have 
the spores and capsules pretty fully developed in the preceding autumn so 
that some of the sporophyte characters are nearly- always accessible. 
Hepatics shrivel more than mosses in drying and are best studied while fresh, 
especially the thalloid forms. 

A few of the rare genera are omitted and in the completed treatment 
some of the minute or difficult species will not be included. 

The Germans call the true mosses Laubmoose, meaning leafy mosses, 
and the Hepatics, Lebermoose, or liver mosses, The name Liverwort was 
originally applied to Marchantia because of its fancied resemblance to the 
liver. Because of this resemblance it was supposed to be a specific for all 
liver troubles according to the old doctrine of signatures. From this came 
the Latin name Hepaticae and the German Lebermoose. "Thus does the 
language of ignorant superstition become the adopted language of science." 
The chief distinctions between Mosses and Hepatics have been noted 
in the Bryologist for April, 1899, but a few additional notes here may prove 

The Hepatics may be leafy stemmed and appear much like mosses, or 
they may consist of a broad, flat and rather thin stem (thallus) which is 
usually closely applied to the substratum. These thalloid Hepatics might 

be mistaken for some of the foliaceous lichens but the Hepatics are always 
much greener and produce spores in a very different manner. 

In the leafy-stemmed Hepatics, often called Scale Mosses, the leaves are 
without midrib and are nearly always in two ranks and flattened so as to lie 
in one plane, but in the great majority of cases there is a third rudimentary 
row on the inner side which are called underleaves, or amphigastra by those 
devoted to technical names. The pedicel which corresponds to the seta of 
the mosses does not, as a rule, grow much until the spores are nearly ripe, 
when it elongates very rapidly. The pedicels and capsules are of a much 
more delicate structure than in the mosses so that they disappear soon after 
the spores have escaped, but the peculiar and characteristic scales or bracts 
around the base of the pedical often remain much longer and help greatly in 
identifying species. Immediately surrounding the base of the pedicel is a 
tubular, somewhat three-sided organ called the inner involucre or perianth, 
surrounding this the outer involucre, called simply involucre by many 
authors. This latter may be either tubular or composed of separate leaf-like 
divisions of varied shapes, called involucral leaves or bracts, or perichatial 
leaves or bracts, or simply bracts. Either one, or even both, of these invo- 
lucres may be lacking in some species. 

So far as possible gametophyte characters have been used in the keys 
and descriptions and in the great majority of cases identification is easy from 
this part of the plant alone. Hepatics generally grow in moist situations on 
soil, roots of trees, and decaying wood. 

Key to Families. 
Plants leafy, mosslike in appearence except for the two-ranked leaves 

entirely lacking midrib Scale Mosses (Jungermanniaceee) . 

Plants consisting of a flattened green thallus, sometimes nearly circular but 
usually elongated and branching. (See illustrations of Riccia, Mar- 

chantia, Anthoceros, etc.) A. 

i. Capsules, if present, immersed in the tissue of the plant. Plants float- 
ing on the surface of still water or floating on the mud along the 

banks Riccia. 

Capsules raised well above the thallus. Plants often growing in mud but 
never floating 2 - 

2. Stomata (in our genera) present, easily discernable with a lens as small 

pores on the upper surface of the rather thick thallus ; capsule borne on 

a special stalked receptacle as in Marchantia. 

Liverworts (Marchantiacea;). 

Stomata not present, on the thinner thallus: capsules never borne on a 

special stalked receptacle 3- 

3. Capsules very long and slender, splitting into two valves when ripe after 
the manner of a mustard pod, the slender hairlike columella remaining 
in the center Horned Liverworts (Anthocerotaceae). 

Capsules globular or ovoid, splitting into four valves; columella lacking. 

Thalloid Scale Mosses (Metzgeriaceag). 

Fig. 4. 

Fig. 6. 

Fig. 5- 
Plate XII. 
Fig. i. a, Sterile and b, fertile thallus Anthoceros punctatus X 2 & 1. 
Fig. 2-6. Marchantia polymorpha, from Bryolouist, 4:34-35, 1901. Fig. 2. 
Male plant a little reduced, showing antheridial receptacles. Fig. 3. 
Longitudinal section of antheridial receptacle magnified. Fig. 4. 
Female plant reduced showing the stalked receptacles which characterize 
this family. These receptacles vary in the family from the shape shown 
in this figure to almost perfectly conical and entire. Fig. 5. Section of 
a part of a female receptacle magnified, showing two sporogonia. The 
seta of one has elongated, pushing the capsule out from the outer fringe 
(involucre) and the inner fringe (perianth) at the base of the seta is a lit" 
tie collar representing the base of the broken calyptra. Fig. 6. Sterile 
thallus with gemma;. 

The plants of this family consist of thallus of medium to large size, one- 
half to six inches in length, usually branching dicotomously but sometimes 
with more than two branches at a fork. They are attached to the substratum 
by numerous roothairs and are thickened in the middle to form a midrib. 
This in some cases is not vary apparent above but shows plainly underneath. 
The upper surface is covered with small pores (stomata) which are very 
apparent with a lens, except in Reboulia. The capsules are spherical or ovoid 


and open irregularly by imperfect valves or by a portion of the top coming 
off after the manner of a lid. In this family the capsules and usually the 
antheridia are borne on special long-stalked receptacles well illustrated by 
the familiar Marchantia. 

Key to the Genera. 
i. Sterile stems bearing abundant gemmae in shallow open receptacles. . . .2 

Sterile stems without gemmae 3 

2 Found only in and around greenhouses: gemmae in crescent-shaped 

receptacles ; never fruiting in our region Lunularia. 

Growing abundantly everywhere; gemmae in cup-shaped receptacles: cap- 
sule-bearing receptacle with 7-11 conspicuous rays Marchantia. 

3. Thallus large ; 2-6 inches long and l / 2 inch or more wide, distinctly areo- 
late as in Marchantia, but areolas larger and hexagonal. . . Conocephalus. 

Thallus less than two inches in length and much narrower 4 

4. Pores (stomata) scarcely distinguishable ; antheridia in sessile receptacles 
which might be mistaken for gemmae; thallus purple on the margins; 
midrib strong underneath but not conspicuous above Reboulia. 

Pores conspicuous, white; antheridia in peduncled disk-like receptacle; thal- 
lus with numerous dark purple scales underneath Preissia. 

Pores conspicuous; antheridia immersed in the thallus; thallus purple 
underneath, at least along the margins 5 

5. Perianth conspicuous, split into 8-16 fringe-like lobes; peduncle not 

chaffy Asterella. 

Perianth lacking: peduncle chaffy at top and bottom Grimaldia. 

(The Reboulia of this key is the Asterella of Gray's Manual and the 
Asterella is the Fimbriaria of that work. ) 


The spore bearing portion of plants of this family is like that of the 
Scale Mosses, but the green part of the plant is a thallus instead of a leafy 
stem in nearly all cases. There are, however, some intermediate forms in 
the family in which the thallus is divided into leaflike lobes. The thallus is 
much less highly differentiated than in the Liverworts and Riccias: there 
are no areolae or pores (stomata), and the thallus is much thinner than in the 
Liverworts, in some species consisting of only a single layer of cells except 
at the midrib. The capsules are borne singly on setae arising directly from 
the thallus. They are spherical to elongated-ovoid and remain enclosed in 
the calyptra until mature when the seta? rapidly elongate and break open the 
calyptra which is left at the base of the seta. The capsules open by four 
valves as in many of the Scale Mosses. A careful search of wet bare earth 
in shaded or springy places will nearly always yield one or more species of 

this family. 

Key to the Genera. 

1. Thallus with a distinct midrib 2. 

Thallus without a distinct midrib 4- 

2. Thallus 1/25 to 1/12 inch wide, dichotomously branched, cilliate along 

the margins Metzgeria. 


Thallus ^ to y 2 inch wide, not cilliate at margins, entire or lobed 3. 

3. Thallus simple or only once forked, 1 to 4 inches long, prostrate: margins 

sinuate to entire: capsule ovoid-cylindric Pallavicinia. 

Thallus dichotomously branched, j^ to 1^ inches long, often densely clus- 
tered and ascending, margins lobed: capsules spherical without perianth, 
^^ appearing buried in the midrib for 

•v. ^i ®wJf£\ sft> some time before the ripening of spores 

^Mtf! W^^^^ML* (Fig- 7) Blasia. 

vw 1 ^a^k6^^^^^ ■*• Thallus pinnately or palmately branch- 

\w?-- n^i/ft; e< ^' J / 24 to I//I2 * n w 'de (except 

)ffy a Fi'WJfr b R. pinguis) Riccardia. 

•^Sg Thallus subsimple or dichotomously 

branched, y& to 'A, inch in width (Fig. 
tlg - 7 - 8) Pellia. 

Fig. 7. Blasia pusilla L. a. Fertile plant in August 
showing capsule in position. At the side is shown the cap- 
sule removed from the thallus. b. Sterile plant with flask- 
shaped bodies which produce gemmae. 

,,,, v . Fig. 8. Pellia epiphylla Raddi. Thallus X 1, showing 

involucre and position of capsule as it appears in August. 
Fig. " 


The reproductive part of the Scale Mosses, including the ripened cap- 
sule and its connected parts, perianth, involucre, etc., is essentially as in 
the Thalloid Scale Mosses, but the vegetative part strongly resembles the 
true mosses in general appearance. The leaves, however, are apparently 
flattened out into two rows, one on either side of the stem. They are 
entirely without midrib and are frequently two-cleft or lobed. One of the 
lobes is often smaller and folded under the other making the leaves " com- 
plicate-bilobed," in the language of the books as shown in the illustrations 
of Radula and Porella. This can best be made out by holding a single 
stem up to the light and examining with a lens, when the under lobe will 
show plainly as a deep shadow. In Scapania, the under lobe is the larger 
and the plants look as if there were four rows of leaves. The lower lobe is 
called the lobule and the upper simply the lobe. Very many species have a 
third rows of leaves on the under side of the stem called technically " amphi- 
gastra" or underleaves, these vary in size from one-third the size of the 
ordinary leaves to so minute that high powers of the compound microscope 
are needed to see them clearly. The upper margin of the leaves may over- 
lap the lower margins of the leaves next above as in Porella, or the upper 
margin of a leaf may lie under the lower margin of the leaf next above as 
in Plagiochila. In the former case the leaves are said to be incubous, in 
the latter succubous. As this distinction is in most cases easy to observe, 
it is given a prominent part in the key. Occasionally the leaves are so far 
apart that it is hard to determine the leaf arrangement, but a careful search 


will usually discover some plants in which this character can be seen. 
In plants with incubous leaves the bud is turned downward: when the 
leaves are succubous the terminal bud is turned up. So far as possible the 
key has been based upon the leafy or vegetative portions of the plants, but 
in some few cases the characters connected with the reproductive organs 
and capsules are necessary to accurately determine a plant. In most cases 
the characters used can be determined without mounting, if, however, they 
can not be readily made out the parts should be mounted as for the com- 
pound microscope. If one has access to a compound microscope itwill often 
prove a very great help, although not necessary to make out the characters 
mentioned. Mnium and Fissidens are sure to be mistaken for Hepatics by 
the beginner unless the midrib or the leaves is noted. 

Key to the Genera and Species. 

i. Leaves entirely or in large part composed of hair-like divisions (easily 

observed if held up towards a strong light) 2. 

Leaves not as above 3. 

2. Plants grayish green, growing over the ground amid mosses in cool 
bogs, at least twice pinnate and somewhat resembling the Fern Mosses; 
leaves divided to base into hair-like lobes Trichocolea. 

Plants dark green, much smaller, growing chiefly on rotton wood, but also 
found on humus-covered stones and soil ; leaves with a considerable 
solid portion Ptilidium. 

Plants exceedingly minute, looking like a small green alga or moss proton- 
ema. Scarcely recognizable except when fruited; common on decayed 
wood, moist soil, etc Blepharostoma. 

3. Leaves incubous A. 

(Scapania and Chiloscyphus forms may be sought here.) 

Leaves succubous B. 


1. Leaves complicate-bilobed, upper lobes entire or nearly so (except 
Jubula). See figures and description of Porella 2. 

Leaves sometimes lobed or cleft but not complicate-bilobed 5. 

2. Plants blackish or brownish green, minute, leafy stems 1/25 inch or less 
wide; lobule like an inflated sac (Plate XIV) Frullania. 

Plants often dark olive-green but not often blackish; 1/16 inch in width, 
lobule not sac-like 3. 

3. Under leaves lacking; perianth strongly flattened crosswise (Plate 
XIII) Radula. 

Underleaves conspicuous 4 . 

4. Lobule with its longer edge attached to lower margin of lobe (See cuts, 
Plate XIII) Lejeunea. 

Lobule with its shorter margin attached to the lower edge of lobe (Plate 
XIV) Porella. 

5. Leaves mostly entire Kantia. 

Leaves strongly toothed, notched, or cleft at apex 6. 

6. Leafy stems less than 1/25 inch in width Lepidozia. 


Leafy stems 1/16 to % inch in width, with downward growing stolons 
(Plate XIII) Bazzania. 


1. Leaves complicate-bilobed, lobes nearly equal or the lower larger giv- 
ing the appearance of four rows of leaves of which the two upper are 
incubous and the two lower succubous Scapania. 

Leaves not complicate-bilobed, in some cases toothed or divided 2. 

2. Leaves undulate on the margin; plants densely clustered; roothairs 
bright claret colored Nardia hyalina. 

Leaves entire or slighly emarginate ; roothairs colorless 3- 

Some or usually all of the leaves strongly toothed or lobed; roothairs color- 
less 7- 

3. Leafy stems at least ]/% inch wide, leaves plainly overlapping, on ground 
and over mosses 5- 

Plants about y% inch wide: many leaves not overlapping 4- 

Leafy stems 1/16 inch wide or less 6. 

4. Aquatic, floating, underleaves absent. 

Chiloscyphus polyanthus var. rivularis. 
On old logs and moist ground, underleaves present Chiloscyphus. 

5. Plants creeping; leaves oblong to oblong-ovate, decurrent — Liochlaena. 
Plants ascending; leaves round obovate, not decurrent Plagiochila. 

6. Leaves with a border of larger cells which appear as a whitish margin 
under the lens Odontoschisma and Nardia crenulata. 

Leaves without border of larger cells Jungermannia Schraderi. 

7. Upper leaves with a strongly many toothed margin Plagiochila. 

Leaves 3-5 cleft Jungermannia barbata. 

Leaves two toothed or cleft 8. 

8. Plants minute, leafy stems less than 1/25 inch wide ; underleaves absent 
or so small as to be invisible with a lens: leaves round-ovate to 

obovate, cleft for at least ]^ their length Cephalozia. 

(Some small species of Jungermannia may be sought here but their 

leaves are less deeply cleft and the plants are a much darker green). 

Leafy stems at least 1/16 inch wide: leaves two toothed but scarcely cleft .9 

9. Underleaves % the size of the other leaves Lophocolea minor. 

Underleaves absent or minute 10. 

10. Leaves subvertical, varying from bidentate to refuse, or even entire 
near apex Lophocolea heterophylla. 

Leaves with the edge attached nearly lengthwise of the stem, extending out at 
almost right angles from it and lying flat in a horizontal plane. Geocalyx. 

Leaves subvertical and all alike inserted more nearly crosswise of the 
stem 11 - 

n. On sterile ground in open woods Jungermannia excisa. 

On rotton wood I2 - 

12. Leaves round-ovate with an obtuse sinus between the teeth. Harpanthus. 

Leaves subrectangular with an acute sinus; plants dark to brownish 
g re en Jungermannia Michauxii. 

Figure 9. 

Figure 10. 

Figure ii. 
Plate XIII. Bazzania, Radula, Lejeunea. 


Fig. 12. 


Fig. 13. 

Fig. 15. 

Fig. 14. 

Plate XIV. Porella, Frullania. 

- 9 8- 


Plate XIII. Fig. 9. Bazzania trilobata (L.) S. F. Gray. From Bryologist, 
4:68, 1901. A. Plant slightly magnified showing flagella springing 
from the underside. B. (1.) Portion of female plant with capsule. (2.) 
Capsule open. C. Involucre, perianth and base of seta enlarged. 
The involucre consists of the small leaves at the bottom of the fig- 
ure. D. Male plant seen from below, showing antheridial branch, 
minute underleaves and incubous arrangement of leaves. E. & F. 
Illustrate spiral elators, spores and cell structure of leaf which cannot 
be seen clearly with a hand-lens. 

Fig. 10. Radula complanata Dumort. A. Plant natural size. B. Branch 
with fruit showing clearly the seta and capsule, with the calyptra at 
base of seta showing through the transparent tubular perianth, and at 
base of the perianth, the involucre. This misrepresents the leaves, 
making them appear succubous. C. Leaf showing lobule with roothairs 
and larger lobe with gemmae along the edge. This illustrates the sim- 
plest forms of " complicate-bilobed " leaf. There are no underleaves. 
D. Calyptra. E. Spores, highly magnified. 

Fig. 11. Various species of Lejeunea, from Bryologist, 6:-27, 1903. 
Showing underleaves in all but the right hand figure. Note that the 
lobule is attached to the lobe by its longer edge. 

Plate XIV. Fig. 12. Porella pinnata L. From the Bryologist, 5: 34, 
1902. A. Underside of stem showing narrow underleaves and narrow 
lobules attached by their shorter edge to lobe. B. Single leaf showing 
lobe and lobule. 

Fig. 13. Porella platyphylla (L.) Lindb. From Bryologist, 5: 35, 1902. 
B. Upper side of stem showing perianth and emerging capsule. Also 
showing clearly incubous arrangement of leaves. C. Underside of 
stem, the leaves shown too far apart. D. Longitudinal section of 
perianth. E. Capsule. F. Leaf. G. Part of plant showing male 

Fig. 14. Frullania. From Bryologist, 5 : 4, 1902. I. Plant of Frullania 
Eboracensis Gottsche., on the bark of birch. II. Underside of same 
showing underleaves and the queer saclike inflated lobules which 
remind one of the bladders of Utricularia. III. and IV. Under and 
upper side of F, Asagrayana Mont. V. Involucre and perianth of F. 

Fig. 15. Ptilidium ciliare (L.) Nees. a. Leaf X 37. b. Plant with 
perianth and young capsule X 2. c. Portion of plant X 5.